Abstract
Drought stress negatively affects plant metabolism and growth, triggering complex defence mechanisms to limit damage. This study evaluated the effectiveness of a single foliar application of 1 mM L-proline (Pro) in winter wheat (Triticum aestivum L., cv. Bohemie) in two separate experiments differing in the time interval between application and drought—7 days (experiment 1) and 35 days (experiment 2). Net photosynthetic rate (A), transpiration rate (E), stomatal conductance (gs), leaf water potential (Ψw), intrinsic water use efficiency (WUEi), endogenous proline content (Pro), malondialdehyde content (MDA), and maximum quantum yield of photosystem II (Fv/Fm) were measured. In experiment 1, drought markedly reduced net photosynthetic rate, transpiration rate, stomatal conductance, and leaf water potential in both drought-stressed treatments, namely, without priming plants (S) and with Pro priming plants (SPro). Pro and MDA content increased under stress. Higher E and gs in the SPro treatment indicated more effective stomatal regulation and a distinct water use strategy. Pro content was significantly lower in SPro compared to S, whereas differences in MDA levels between these treatments were not statistically significant. The second drought period (D2) led to more pronounced limitations in gas exchange in both S and SPro. Enhanced osmoregulation was reflected by lower Ψw (S < SPro) and higher Pro accumulation in S (S > SPro). The effect of exogenous Pro persisted in the form of reduced endogenous Pro synthesis and improved photosystem II protection. Rehydration of stressed plants restored all monitored physiological parameters, and Pro-treated plants exhibited a more efficient recovery of gas exchange. Experiment 2 demonstrated a long-lasting priming effect that improved the preparedness of plants for future drought events. In the SPro treatment, this stress memory supported more efficient osmoregulation, reduced lipid peroxidation, improved protection of photosystem II integrity, and a more effective restart of gas exchange following rehydration. Our findings highlight the potential of exogenous proline as a practical tool for enhancing crop resilience to climate-induced drought stress.
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Claims 18
Claim · model-written summary
Drought reduced photosynthesis, transpiration and stomatal conductance in both stress groups.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- droughte0
- Outcome
- net photosynthetic rate, transpiration rate, stomatal conductancee0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“drought markedly reduced net photosynthetic rate, transpiration rate, stomatal conductance, and leaf water potential in both drought-stressed treatments”
t0“Wheat”
Claim · model-written summary
Drought reduced leaf water potential in both stress groups.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- droughte0
- Outcome
- leaf water potentiale0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“drought markedly reduced net photosynthetic rate, transpiration rate, stomatal conductance, and leaf water potential in both drought-stressed treatments”
t0“Wheat”
Claim · model-written summary
Proline and MDA content increased under stress.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- stresse0
- Outcome
- Pro and MDA contente0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Pro and MDA content increased under stress”
t0“Wheat”
Claim · model-written summary
Primed stressed plants had higher transpiration and stomatal conductance.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- SPro treatmente0
- Outcome
- E and gse0
- Direction
- Highere0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Higher E and gs in the SPro treatment”
t0“Wheat”
Claim · model-written summary
Proline content was significantly lower in primed than unprimed stressed plants.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Pro content was significantly lower in SPro compared to S”
t0“Wheat”
Claim · model-written summary
MDA differed nonsignificantly between primed and unprimed stressed plants.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- these treatmentse0
- Outcome
- MDA levelse0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“differences in MDA levels between these treatments were not statistically significant”
t0“Wheat”
Claim · model-written summary
The second drought period caused more pronounced gas exchange limitations in both groups.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- second drought period (D2)e0
- Outcome
- limitations in gas exchangee0
- Direction
- more pronouncede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The second drought period (D2) led to more pronounced limitations in gas exchange in both S and SPro”
t0“Wheat”
Claim · model-written summary
Unprimed stressed plants had lower water potential than primed stressed plants.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“lower Ψw (S < SPro)”
t0“Wheat”
Claim · model-written summary
Unprimed stressed plants accumulated more proline than primed stressed plants.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“higher Pro accumulation in S (S > SPro)”
t0“Wheat”
Claim · model-written summary
Exogenous proline reduced endogenous proline synthesis.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- exogenous Proe0
- Outcome
- endogenous Pro synthesise0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“effect of exogenous Pro persisted in the form of reduced endogenous Pro synthesis”
t0“Wheat”
Claim · model-written summary
Exogenous proline improved photosystem II protection.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- exogenous Proe0
- Outcome
- photosystem II protectione0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“exogenous Pro persisted in the form of reduced endogenous Pro synthesis and improved photosystem II protection”
t0“Wheat”
Claim · model-written summary
Rehydration restored all monitored physiological parameters.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Rehydratione0
- Outcome
- all monitored physiological parameterse0
- Direction
- restorede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Rehydration of stressed plants restored all monitored physiological parameters”
t0“Wheat”
Claim · model-written summary
Proline treated plants recovered gas exchange more efficiently.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Pro-treatede0
- Outcome
- recovery of gas exchangee0
- Direction
- more efficiente0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Pro-treated plants exhibited a more efficient recovery of gas exchange”
t0“Wheat”
Claim · model-written summary
Long interval priming improved preparedness for later drought.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- priminge0
- Outcome
- preparedness of plants for future drought eventse0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Experiment 2 demonstrated a long-lasting priming effect that improved the preparedness of plants for future drought events”
t0“Wheat”
Claim · model-written summary
Stress memory supported more efficient osmoregulation.
- Taxon
- wheat
Triticum aestivumt0 - Outcome
- osmoregulatione0
- Direction
- more efficiente0
- Qualifier
- stress memorye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“this stress memory supported more efficient osmoregulation”
t0“Wheat”
Claim · model-written summary
Stress memory reduced lipid peroxidation.
- Taxon
- wheat
Triticum aestivumt0 - Outcome
- lipid peroxidatione0
- Direction
- reducede0
- Qualifier
- stress memorye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“this stress memory supported more efficient osmoregulation, reduced lipid peroxidation”
t0“Wheat”
Claim · model-written summary
Stress memory improved photosystem II integrity protection.
- Taxon
- wheat
Triticum aestivumt0 - Outcome
- protection of photosystem II integritye0
- Direction
- improvede0
- Qualifier
- stress memorye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“this stress memory supported more efficient osmoregulation, reduced lipid peroxidation, improved protection of photosystem II integrity”
t0“Wheat”
Claim · model-written summary
Stress memory supported more effective gas exchange restart after rehydration.
- Taxon
- wheat
Triticum aestivumt0 - Outcome
- restart of gas exchangee0
- Direction
- more effectivee0
- Qualifier
- stress memorye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“this stress memory supported more efficient osmoregulation, reduced lipid peroxidation, improved protection of photosystem II integrity, and a more effective restart of gas exchange following rehydration”
t0“Wheat”
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